AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 100:89-100 (1996)

Sexual Dimorphism in the Pelvic Midplane and Its Relationship

to Neandertal Reproductive Patterns
DANA E. WALRATH AND MICHELLE M. GLANTZ
Department of Anthropology, University of Pennsylvania, Philadelphia,
Pennsylvania 19104

KEY WORDS Birth canal, Bispinous diameter, Computed

tomography, Neandertal reproduction, Sexual dimorphism

ABSTRACT The fragmentary nature of the fossil record has limited the
analysis of the Neandertal pelvis to the superior pubic ramus and the pelvic
inlet. From a n obstetric viewpoint, the pelvic midplane or “plane of least
dimensions,” defined by the distance between the ischial spines, must be
considered in the analysis of hominid reproduction. We examined the relation-
ship between BSD and weight in a mixed sex hospital population undergoing
diagnostic computed tomography (CT) scans (41 females and 40 males). Be-
cause femoral head diameter squared (FH2) has been used a s a proxy for
weight in skeletal populations, it was also analyzed with respect to BSD and
weight. Bivariate regression analysis of BSD with other body dimensions
indicates the presence of significant sex differences. In females, but not in
males, weight is a statistically significant predictor of BSD. FH2 is a n even
better predictor of BSD in females while nonsignificant in males. Although
weight and FH2are significantly correlated with BSD in females, FH2does not
predict weight in females as well a s it does in males. The positive correlation
between skeletal frame size and BSD in females is indicative of a n evolution-
ary pattern that must take into account the pressures of reproduction. Our
results indicate that critical dimensions of the pelvis must increase a s the
maternal skeleton becomes larger. These results provide a context for the
interpretation of the reproductive patterns of a relatively robust hominid
population like the Neandertals. o 1996 Wiley-Liss, Inc.

The observation that the Neandertal supe-
rior pubic ramus is elongated and thin in
cross-section (McCown and Keith, 1939;
Stewart, 1960; Smith, 1976; Trinkaus, 1976)
compared to that of modern humans has led
to the construction of a number of theories
concerning Neandertal reproductive pat-
terns (Anderson, 1989; Brothwell, 1975;
Dean et al., 1986; Greene and Sibley, 1986;
Rosenberg, 1988; Trinkaus, 1984,1990; Wol-
poff, 1980). Wolpoff (1980) proposed that a
longer superior pubic ramus contributed to
a larger pelvic inlet which, in turn, corres-
ponded to a relatively larger neonate cra-
nium. Subsequent interpretations, based on
the morphology of the Neandertal pelvis,
have included the prolonged gestation
length model (Trinkaus, 1984) and the accel-
erated fetal growth model (Dean e t al., 1986).
The most recent contribution to these
functional interpretations of Neandertal pel-
vic morphology is Rosenberg’s maternal
weight model (1988). Building on the finding
that maternal body weight is a critical deter-
minant of birth weight (Garn and Pesick,
19821, Rosenberg examined the relationship
between maternal body weight and pelvic
Received January 18, 1995; accepted November 7, 1995.
Address reprint requests to Dana Walrath, Department of An-
thropology, University of Pennsylvania, Philadelphia, PA 19104.

0 1996 WILEY-LISS, INC

90 D.E.WALRATH AND M.M.GLANTZ
Fig. 1. Plane of least dimensions in the obstetric pelvis. (Reproduced from Oxorn-Foote, 1986, with
permission of Appleton-Century-Crofts, Norwalk, CT.)
inlet dimensions. Her data suggested that
populations with the highest relative
weight, like the Zuni, possess the longest
pubic bones. Although no modern human
population has pubic bone dimensions that
match those of Neandertals, it is unlikely
that any modern human population is as
heavy relative to their stature as the Nean-
dertals. Therefore, Rosenberg suggested
that Neandertal infants were the same size
at birth relative to their mothers as modern
infants. Her model linked superior pubic ra-
mus length and birth weight to the absolute
and relative weight of the mother. Conse-
quently, Rosenberg concluded that the repro-
ductive patterns of Neandertals and modern
humans are similar.
The fragmentary nature of the fossil re-
cord has dictated that interpretations of the
Neandertal pelvis and reproductive patterns
rely solely on the pelvic inlet and its corres-
ponding structures. However, the emphasis
on the pelvic inlet in these reconstructions is
of limited use from a n obstetric perspective.
Although the pelvic inlet becomes contracted
in response to nutritional stress during de-
velopment, it is the pelvic midplane (Fig. l ) ,
the narrowest point in the birth canal, that
interferes with the birth process in most pop-
ulations (Sibley et al., 1992; Oxorn-Foote,
1986; Floberg et al., 1986, 1987). The dis-
tance between the ischial spines, or bispi-
nous diameter (BSD), defines the obstetric
“plane of least dimensions” (Oxorn-Foote,
1986). Projecting into the birth canal, the
ischial spines provide support for the anal
sphincter and for the abdominopelvic organs
to resist intraabdominal pressure. The mus-
cles and fascia of the pelvic floor insert di-
rectly or indirectly onto the ischial spines.
Thus, according to Abitbol, “the ischial
spines in humans pose the most serious
threat to obstetrical delivery of any single
feature of the human pelvis” (1988:58).
Unfortunately, ischial spines are not pre-
served in the Neandertal pelvic samples. The
only reasonably complete Neandertal pelvis
(Kebara 2) lacks ischial spines and is a male
(Rak, 1990; Rak and Arensburg, 1987;
Tague, 1992). Further, the appropriateness
of attributing this specimen to the Neander-
tals recently has been the subject of debate
(Arensburg, personal communication). In
the absence of completely preserved Nean-
dertal ischial spines, modern human data
provide a context through which Neandertal
pelvic relationships may be inferred.
To provide a reference for the interpreta-
tion of Neandertal pelvic morphology, we ex-
amine the pelvic midplane (BSD) in a mod-
 SEXUAL DIMORPHISM IN THE PELVIC MIDPLANE
TABLE 1. SamDle
n 41
Age range (average)
Weight range (average) in kilograms
Height* range (average) in centimeters
Ethnicity
'For height, n = 27 females and 23 males.
ern sample. BSD constrains the descent of
the fetus through the pelvis during birth.
Thus, birth of large neonates should require
a correspondingly large maternal BSD. Al-
though BSD has been analyzed extensively
in the obstetric literature in terms of its con-
tribution to fetal-pelvic disproportion (e.g.,
Jagani et al., 1981; Oxorn-Foote, 1986;
Morgan e t al., 1986; Thurnau and Morgan,
1992), the relationship of the pelvic mid-
plane to maternal anthropometrics has not
been systematically investigated. Because
maternal weight is a strong predictor of birth
weight (Garn and Pesick, 1982), we propose
that selective forces acting on the female pel-
vis would favor a strong correlation between
BSD and weight in females. To test a gener-
alized maternal size-pelvic midplane hy-
pothesis and to examine whether a particu-
lar component of maternal weight (i.e., bone
mass, metabolic stores, height) may underlie
this correlation, we analyzed the relation-
ship between BSD and the following anthro-
pometric measures: weight, height, body
mass index (BMI), and skeletal frame size.
Femoral head diameter squared (FH2)is
used in this study a s a proxy for skeletal
frame size. Because i t is the articular surface
of a weight-bearing joint, many authors have
used the femoral head as a n index of body
size in skeletal specimens (McHenry, 1976;
Jungers, 1988; Rosenberg, 1988). Ruff and
colleagues (1991) have shown that the femo-
ral head remains constant relative to femo-
ral cortices despite lifetime weight fluctua-
tions. Thus, the femoral head can be of use
in distinguishing soft vs. bony components
of weight in the mixed age sample of known
weights. Skeletal frame size is not a measure
of bone density. We propose that larger val-
ues of FH2are indicative of overall bone mas-
91
Females Males
40
23-72 (47) 24-70 (49)
41-104 (64) 57-118 (82)
152-175 (162) 165-188 (178)
54% European 49% European
10% African-American 7% African-American
3% Asian 44% unknown
33% unknown
siveness, while smaller values indicate rela-
tive gracility or slightness of the skeletal
frame.
Because the pelvic inlet has been the focus
in reconstructions of Neandertal reproduc-
tive patterns (Anderson, 1989; Brothwell,
1975; Dean et al., 1986; Greene and Sibley,
1986; Rosenberg, 1988; Trinkaus, 1984,
1990; Wolpoff, 1980), we also analyzed the
relationship between the pelvic inlet and
midplane. Males serve as a reference for non-
specific effects of body size on pelvic geome-
try, as they are not directly subject to selec-
tive pressures imposed by parturition.
METHODS
We measured pelves from computed to-
mography (CT) scans from 41 females and
40 males undergoing diagnostic studies at
the Hospital of the University of Pennsylva-
nia (Table 1). Subjects ranged in age from
23-72 years with a mean of 48 years. CTs
were taken on the subjects for a variety of
reasons including trauma and for the diag-
nosis of specific disease states. All subjects
included in the sample have fused bones
lacking pathologies or age-related degenera-
tive changes. The following exclusion crite-
ria were used to determine a n appropriate
study sample: individuals indicating that
they had experienced weight loss or gain of
20 pounds or more in the past 6 months,
those who were extremely obese or lean as
defined by the ninety-fifth and fifth centiles
from the National Health and Nutrition Ex-
amination Survey (NHANES) standards
(Frisancho, 1990), and patients identified a s
having end-stage illness.
Before measurements were taken, the fol-
lowing methods were explored to ascertain
92 D.E. WALRATH AND M.M. GLANTZ

Fig. 2. Scout CT scan of pelvis illustrating (1)bispinous diameter, distance between the ischial spines
(BSD), and (2) femoral head diameter, maximum diameter of the femoral head (FH).

the optimal technique for measurement of sired distance interval between axial sec-
the pelvic midplane. Plane anteroposterior tions necessary to image specific structures
X-ray was found to be unsuitable for this in cross-section. However, with regard to the
study because of magnification error. The pelvic midplane, the position of the subject
distance from the X-ray source to the ischial affects the probability of capturing both of
spines to the film is variable depending on the ischial spines in a single axial section.
the amount of soft tissue in the gluteal re- Even if the spines are present in a single
gion of the subject. Because of this variable section, it is difficult to determine which por-
factor, the calculation of case-specific magni- tion of the spine is observed (Aronsen and
fication could not be accomplished without Kier, 1991). Because it is only the most distal
exposing the subjects to additional lateral portions of the spines that are measured
X-rays. Ultrasound technology also was when deriving the BSD, scout CTs instead
found to be unsuitable because the ischial of axial sections were used for measurement
spines were not visible. in this study.
Computed tomography proved to be the The accuracy of the measurements taken
most reliable method for measuring the pel- from scout films was verified by the compari-
vic midplane (Fig. 2). Because the subject son of these same measurements taken from
is exposed to radiation from a 360 degree a modern bony pelvis with calipers (Fig. 3).
radius, it is possible to reconstruct mathe- Measurement error was less than 1%for
matically a n anatomically accurate image each value. In addition to BSD, we measured
of the skeleton and other tissue structures acetabulo-symphyseal length (PAC) and
(Thurnau and Morgan, 1992). “Scout” films transverse diameter (TD) of the pelvic inlet
resembling conventional anteroposterior (Table 2). PAC was measured from the most
X-ray images are used to determine the de- superio-medial aspect of the pubic symphy-
 SEXUAL DIMORPHISM IN THE PELVIC MIDPLANE 93

Fig. 3. Transverse diameter (l), the maximum distance of the pelvic inlet from a medio-lateral
perspective (TDj; bispinous diameter (2) (BSDj; and acetabulo-symphyseal length (3j, the most superio-
medial aspect of the pubic symphysis to the most superior aspect of the acetabulum (PAC).

TABLE 2. Ranges and averages of pelvic and femoral head measurements in millimeters
F ema 1es Males
Bispinous diameter range (average) 83-112 (100) 68-94 (80)
Transverse diameter of pelvic inlet range (average) 111-147 (128) 95-137 (113)
Acetabulo-symphyseal length range (average) 92-126 (107) 93-120 (108)
Femoral head range (average) 3 6 4 8 (41) 42-55 (48)

sis to the most superior aspect of the acetab-
ulum. This CT approach precluded measure-
ment of the pubis from its termination in
the acetabulum, used in classic studies of
pelvic morphology (Washburn, 1948, 1949;
Hanna and Washburn, 19531, but proved
highly reproducible. In this regard, the PAC
values derived in this study are not directly
comparable to the published Neandertal pu-
bic bone values because the acetabulum is
missing in many of these specimens (Trin-
kaus, 1984; Rosenberg, 1988). TD is defined
as the maximum distance of the pelvic inlet
from a medio-lateral perspective.
To quantify skeletal frame size, we mea-
sured the maximum diameter of the femoral
head using the technique of Ruff and col-
leagues (1991). Following Rosenberg (19881,
this value was squared (FH2)as a n index of
the area of this weight-bearing joint. Be-
cause weight was recorded for all 8 1 of our
subjects, we also tested the utility of femoral
head a s a proxy for body weight. In addition,
we tested the potential differences in the
ability of FH, FH2, and FH3 to predict body
weight in order to compare the power of this
measure as a diameter, a n area, and a vol-
ume. All skeletal measures were taken three
times and then averaged.
In addition to these skeletal measures, we
derived the body mass index (BMI) by divid-
ing weight in kilograms by height in centi-
meters squared for those individuals whose
height was recorded (27 females, 23 males).
RESULTS
Bivariate regressions of BSD with other
body dimensions are reported in Table 3. Ab-
solute weight is a significant predictor of
BSD only in females ( P = .01, R2 = .15).Nei-
ther relative weight, as defined by BMI, nor
height serve as significant predictors of BSD
in either sex. However, in females P values
for regressions of BSD with height and BMI
approach significance.
Skeletal frame size, represented by FH',
is the most powerful independent predictor
94 D.E. WALRATH AND M.M. GLANTZ

TABLE 3. Biuariate repressions of bisuinous diameter with other body dimensions

Females MaIes
R' P R' P
Weight (kilograms) .1503 .0123* ,0445 .1973
Height (centimeters) .1393 ,0552 .1350 .0925
Body mass index (weightheight') .1315 ,0630 ,0077 ,6982
(Femoral head diameter)' (millimeters)' 5289 <.0001** .0498 ,1722
Transverse diameter (millimeters) ,4453 <.0001** ,1744 .0242*
Acetabulo-svmDhvsed len&h (millimeters) ,4962 <.0001** .1622 .0111*
*P < .05.
**P < .01.

120

9 0 0
0
0

110 0

'D
L
a, 0 -3

/
w 0 0 0
a, 0
0 0 - 0
-ti 100 4 0 0
73
0
cn
3
0
c
-d
0. 4 ' 0 0
/

2 90
n O
I
I /
0
/

80
I I I 1 1 1 1
1200 1400 1600 1800 2000 2200 2400
Femoral head diameter squared
Fig. 4. Bivariate regression of femoral head diameter squared (FH2)and bispinous diameter (BSD).
I n females, a highly significant linear relationship exists between FH2and BSD ( P < .001, R2 = .53).FH2
explains 10%more of t h e variation in BSD when a curvilinear regression with a third-order polynomial is
used (R2 = .58).

of BSD in females ( P < .001, R2 = 5 3 ) . Con-
versely, no relationship exists between BSD
and FH2 in males ( P = .17, R2 = .05).
The relationship between FH2 and BSD
for females and males is graphically illus-
trated in Figures 4 and 5 . These figures show
a positive linear relationship between FH2
and BSD only in female pelves. Moreover,
a third-order polynomial best describes this
relationship in that it comprises the reduc-
tion in variability in BSD values at extreme
ends of the FH2 range. The R2 value for the
curvilinear model is 10%more powerful than
the linear model (R2 = .58 vs. 5 3 ) .
In contrast to the female pattern, male
BSD values are randomly distributed across
the entire FH2 range. Accordingly, the re-
gression line shown in Figure 5 does not sig-
 SEXUAL DIMORPHISM IN THE PELVIC MIDPLANE 95

100

0
0

0
90 0
0 0

L 0 0 0
0, 0 0 0 0
-v
0,
E
(0
-I+ 80
-0 0 0 0
0
cn 0 0
3 0 0
0 0 0 0
c 0
-I+
a 0 0
cn
.A
n 70
0

60
I I I I I I
17b0 19bO 2100 2300 2500 2700 2900 3100
femoral head diameter squared

Fig. 5. Bivariate regression of femoral head diameter squared (FH2) and bispinous diameter (BSD)
in males indicating no association.

nificantly characterize the relationship be- In Table 4,bivariate regressions of weight

tween FH2 and BSD in males.
The distributions of residuals from the bi-
variate regressions of BSD and FH2in males
and females are presented in Figure 6. Fe-
male BSD variation is limited to the middle
values of FH2. In contrast, male BSD varies
throughout the entire range of FH2 values.
In other words, in contrast to females, the
dimensions of male BSD may vary even at
the extreme ends of the FH2 range.
Acetabulo-symphyseal length (PAC) and
transverse diameter (TD) of the pelvic inlet
are significant predictors of BSD in both
sexes (Table 3). Yet both of these skeletal
measures account for nearly three times the
amount of variance in BSD in females com-
pared to that of males. Male BSD deviates
from the pattern of increasing pelvic inlet
dimensions produced by general scaling ef-
fects, while female BSD adheres closely t o
this pattern.
and FH2 with pelvic inlet dimensions are
reported. Weight and FH2 correlate with
PAC and TD at statistically significant levels
in both sexes ( P < .01). By contrast, the sig-
nificant association between the size vari-
ables, weight, and FH2 and the pelvic mid-
plane is found only in females (Table 3).
The ability of femoral head diameter (FH)
to predict body weight was tested. Although
Rosenberg (1988) employed the area of a
weight-bearing joint (FH2)in her study, she
suggested that FH3may more accurately in-
dex body weight because weight reflects a
volumetric measure. Bivariate regressions
of known body weight with FH, FH2,and FH3
are presented (Table 5). Our results indicate
that no significant difference exists in the
ability of any of these femoral head values
to predict body weight. In all three cases,
the femoral head is nearly three times more
powerful for predicting weight in males than
96 D.E. WALRATH AND M.M. GLANTZ

Males

Females

Fig. 6. The distribution of residuals from regressions of BSD predicted by FHZ demonstrate sex
specific patterns. In females, residual distribution is entirely limited to the middle of the FH' range.

females. Male values are highly significant

TABLE 4. Bivariate regressions of weight and (femoral a t the P < .01 level, while female results are
head diameter)' with pelvic inlet dimensions
significant at only the <.05 level.
Females Males
R' P R2 P DISCUSSION
Weight Because of the limitations of the fossil re-
Transverse diameter .2764 ,0004 ,2275 ,0077 cord, reconstructions of Neandertal repro-
Acetabulo-symphyseal ,1652 ,0084 ,2869 .0004
length ductive patterns rely on the dimensions of
(Femoral head diameterY the superior pubic ramus and inferred pelvic
Transverse diameter ,3876 <.001 ,3179 ,0012 inlet size. We redirect the emphasis of these
Acetabulo-symphyseal ,4785 1 . 0 0 1 ,4654 <.001
length reconstructions to the midplane or plane of
least dimensions. This study identifies the
pelvic midplane, reflected in BSD, as a locus
of sexual dimorphism resulting from the se-
TABLE 5. Bivariate regressions of body weight with lective pressure of parturition on female pel-
femoral head diametec (femoral head diameter)', and vic anatomy.
(femoral head diameteri3
Females Males
Pelvic inlet vs. pelvic midplane
R2 P R2 P In the present study, we address the im-
Femoral head diameter ,1420 ,0165 ,4084 <.0001 pact of weight and skeletal frame size on the
(Femoral head diameter)' ,1429 ,0162 ,3977 <.0001 dimensions of the pelvic midplane and inlet.
(Femoral head diameter)' ,1437 ,0159 ,3858 <.0001 BSD is the only pelvic measurement affected
 SEXUAL DIMORPHISM IN THE PELVIC MIDPLANE
by weight and skeletal frame size (FH2)in
a gender-specific manner. In contrast, the
pelvic inlet of both males and females, repre-
sented by PAC and TD, responds similarly
to increases in weight and FH2. I t appears
that the relationship between these inlet di-
mensions and either weight or FH2 reflects
scaling effects that impact both sexes
equally. In contrast, gender-specific relation-
ships between BSD with both weight and
FH2provide evidence for selective pressures
of parturition acting on female pelvic geome-
try. These results support previous studies
which highlight the importance of the pelvic
midplane, rather than the inlet, in recon-
structions of reproductive patterns (Greene
and Sibley, 1986; Abitbol, 1988; Tague,
1992).
In the reconstruction of Neandertal repro-
ductive patterns, the complexity of the rela-
tionship between pelvic geometry and both
reproduction and bipedalism must be consid-
ered. The length of the superior pubic ramus
and its contribution to the dimensions of the
pelvic inlet in Neandertals may have been
associated more strongly with the pressures
of locomotion, posture, (Rak and Arensburg,
1987)or overall body size (Greene and Sibley,
1986; Frayer, 1988) rather than with the re-
quirements of reproduction (Wolpoff, 1980;
Trinkaus, 1984; Dean et al., 1986; Rosen-
berg, 1988).
The presence of a n elongated superior pu-
bic ramus in both Neandertal sexes and even
in the juvenile specimen La Ferrassie 6 led
many researchers to consider these traits to
be genetically determined (Frayer, 1988) and
unique to the Neandertals, given their pro-
posed reproductive adaptation (Tompkins
and Trinkaus, 1987). However, the earlier
hominid specimens with preserved pubic
bones, namely a n A. afarensis, Al-288-1, and
a n A. africanus, Sts-14, also show elongated
superior pubic rami as well a s a somewhat
platypelloid pelvic inlet (Lovejoy et al., 1973;
Tague and Lovejoy, 1986). The interpreta-
tion of the pelvic morphology of Sts-14 is
less straightforward than that of Al-288-1
because its left pubic bone is distorted (Day,
1973). Although the sample size consists of
two specimens from distinct species, Frayer
(1988) suggested that the australopithecines
may have had long pubes in spite of the fact
97
that they did not give birth to big-brained
babies. The only other specimen with par-
tially complete pelvic morphology, WT-
15000, a H. erectus, has been described as
having a large transverse diameter of the
pelvic inlet (Walker and Ruff, 1993).Though
the inlet has been reconstructed without the
benefit of preserved pubes, the inferred pres-
ence of elongated superior pubic rami in this
juvenile male specimen may be interpreted
as a plesiomorphy and have little to do with
increasing neonate cranial dimensions
through time (Abitbol, 1991; Frayer, 1988).
Although the pelvic inlet and midplane are
both dimensions of the birth canal, a reinter-
pretation of the fossil record with respect to
the results of the present study suggests that
the pelvic midplane rather than the inlet
is the critical dimension for successful birth.
Absolute weight vs. skeletal frame size
Our results indicate that skeletal frame
size, rather than weight, has a greater im-
pact on the dimensions of the female pelvic
midplane. Skeletal frame size a s indexed by
FH2 is a more powerful predictor of BSD in
females than height or weight. The emphasis
on skeletal frame size rather than weight
represents a refinement of Rosenberg's ma-
ternal weight model (1988). By focusing on
one of the constituents of body weight, this
study is able to address more directly the
relationship between skeletal size and bony
constraints of the pelvis. Given our results,
maternal skeletal frame size appears to be
a critical factor in determining the female
pelvic dimensions that constrain the birth
process.
The distribution of BSD values for specific
measures of FH2 show a female-specific pat-
tern that is suggestive of stabilizing selec-
tion. Until more data are gathered, it ap-
pears that the female pattern is defined by
random variation in BSD around the middle
portion of the FH2range, whereas variability
is reduced at the extreme ends of this range.
Although the data presented in this study
do not address the relationship between
skeletal frame size and neonate size, they
indicate that skeletally large women must
have a correspondingly enlarged pelvic mid-
plane. It may be inferred that a n enlarged
BSD is related to overall larger neonate sizes
98 D.E. WALRATH AND M.M. GLANTZ
for skeletally large and relatively heavier
females.
The male pelvis is not directly exposed to
the selective pressure of parturition. Conse-
quently, male BSD variability is not con-
strained by specific values of FH2.Male skel-
etal frame size and the dimensions of the
pelvic midplane are unrelated. Male ischial
spines may increase in size and change in
orientation without any compensatory ad-
justments in order to prevent a bony im-
pingement into the birth canal.
Application to the fossil record
The finding that FH2 is a powerful pre-
dictor of female BSD is particularly useful
in the context of the fossil record where body
weight is unknown for even the most com-
plete skeletal remains. In skeletal popula-
tions, weight traditionally has been esti-
mated by some bony parameter such a s the
femoral head diameter (McHenry, 1976;
Jungers, 1988; Rosenberg, 1988).As weights
were known for all subjects in this study,
we tested the reliability of predicting body
weight from femoral head diameter. Bivari-
ate regressions of weight with femoral head
diameter indicate that this estimate is more
reliable in males than in females (Table 4).
This discrepancy between the sexes may be
accounted for by the differences in the con-
stituents of weight for males and females
(Ruff et al., 1991).These data show that skel-
etal frame size rather than body weight dic-
tates pelvic midplane dimensions in females.
The ability to rely solely on bony measures in
the reconstruction of reproductive processes
minimizes the potential error inherent in
body weight inferences from skeletal ma-
terial.
Because FH2values of Neandertal females
with associated pelvic remains (Rosenberg,
1988) fit within the range of our female sam-
ple, these data are particularly relevant to
the reproductive patterns of the Neander-
tals. The success with which Neandertal FH2
predicts BSD is even more likely given the
reduced levels of variability at the extreme
ends of the range in the present study and
the observation that articular surface size
has remained relatively constant through
evolutionary time (Ruff e t al., 1993). Using
the regression equations generated with the
femoral head data presented here, BSD for
these Neandertal specimens would fall
within the upper portion of the modern hu-
man range. Female Neandertal femoral
heads with associated pelvic remains such
as Krapina 208 (FH = 47 mm) and Tabun
C 1 (FH = 43 mm) (Trinkaus, 1984) predict
BSD values of 108 and 102, respectively.
In this study, we established the similarity
between male and female pelvic inlet mor-
phology in its response to absolute and rela-
tive weight and skeletal frame size. We have
also determined that superior pubic ramus
length and transverse diameter explain
nearly three times as much of the variability
in female BSD compared with that of males.
Given these observations from a modern hu-
man sample, we suggest that the elongated
superior pubic ramus in the Neandertals
may be interpreted a s a compensatory ad-
justment to provide adequate BSD in fe-
males from a skeletally large and heavy pop-
ulation.
CONCLUSIONS
We suggest that knowledge of pelvic adap-
tation to parturition in modern humans may
facilitate the interpretation of the reproduc-
tive patterns and pelvic morphology of Nean-
dertals. Two observations pertinent to pelvic
structural adaptation to birth are 1)the posi-
tive correlation between maternal and fetal
size and 2) the obstetric significance of the
pelvic midplane or BSD as the narrowest
point in the birth canal. The female-specific
correlation between BSD and body size is
consistent with the reproductive function of
the female pelvis. In contrast, male BSD var-
ies widely with changes in overall body size.
Although weight has a significant impact on
the female pelvic midplane, BSD is more
highly correlated with maternal skeletal size
as indexed by FH2.In addition, FH2is a bet-
ter predictor of weight in males than in fe-
males, indicating that the use of FH2to esti-
mate female body weight from skeletal
samples may be unreliable. However, i t ap-
pears that relative skeletal frame size has
a greater impact on bony interrelationships
and dimensions in the pelvis than does ma-
ternal weight, thereby reducing the impor-
tance of constructing reliable weight estima-
 SEXUAL DIMORPHISM IN THE PELVIC MIDPLANE
tions. The following are two issues raised by
this study: 1)is relative skeletal frame size
a better predictor of neonate size than
weight?, and 2) might a similar relationship
between FH2and BSD hold for Neandertal or
other skeletal remains? Until more complete
fossil evidence becomes available, estimates
of Neandertal female BSD may be derived
from FH2values. Application of the skeletal
frame size model may provide new insights
into the Neandertal reproductive process.
ACKNOWLEDGMENTS
The authors acknowledge the constructive
advice and critical comments of Peter Bing-
ham, George Boyajian, Robert S.O. Hard-
ing, Rebecca Huss-Ashmore, Francis Johns-
ton, Michelle Lampl, Alan Mann, Janet
Monge, and Phillip Tobias. We would also
like to thank Bernard Birnbaum, Gabor Her-
mann, Morrie E. Kricun, and the computed
tomography technical staff at the Depart-
ment of Radiology, Hospital of the University
of Pennsylvania. Anonymous reviewers im-
proved the final draft of this work. This work
was supported in part by a National Science
Foundation graduate fellowship to the
first author.
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